Population Estimation Procedures Absolute versus Relative Density Absolute Versus Relative Density Catch Per Unit Effort Nearest Neighbour Estimates Transect Sampling Hayne’s Estimator Line Transect Estimators Sightability Models Mark-Recapture Techniques Examples of combined applications Absolute Density Estimates Number per unit area Relative Density Estimates Number per unit effort Catch Per Unit Effort Catch Per Unit Effort Catch Per Unit Effort Nearest Neighbour Estimates 1 What if detection changes with distance? Transect Sampling Density = N / L x 2W Length (L) W W Hayne’s Estimator Hayne’s Estimator White-eared Kob Data (after Krebs 1989) Line Transect Estimators 2 Line Transect Estimators Line Transect Estimators g(X) 1 w Line Transect Estimators Line Transect Estimators This image cannot currently be display ed. This image cannot currently be display ed. g(X) 1 w Line Transect Estimators This image cannot currently be display ed. Assumptions Distance sampling requires that: all animals on or near the transect line must be seen, distances from the transect to the animals are measured accurately, and animals are spotted and accurately located before they move in response to the approaching observer. 3 Sightability Models Advantages Each each population estimate requires only one sample or overflight Natural movements and double counting are not a problem Technique is robust to variable visibility of animals within and between surveys Technique, sampling protocols, and statistics have been extensively developed. Sightability Models Visibility of an animal depends in part on the habitat is in, group size, the observer, animal activity, and perhaps even the sex of the animal. Sightability Models Sightability Models: Assumptions Groups are sighted independently and that all meaningful covariates affecting sighting rates are included in the model Model development requires an adequate sample size (100-200 groups) Technique may not be the best choice for ungulates in situations where detection rates are low, especially when detection rates average less than 50%. Model developed based on the sightability rate of marked or ground-truthed animals Model predicts sighting probability of individual groups of animals based on a set of covariates such as group size, percent tree and shrub cover that will hide animals, percent snow cover, observer experience, survey intensity, etc. The logistic regression model of the sighting covariates is then applied to the sighting conditions recorded for each count unit and each entire survey. Advantages and disadvantages Sightability models are most useful if sighting rates for all groups are >60% Models are specific to covariates affecting sighting rates; hence, different model variants are required for different aircraft, different seasons, and different landscapes Considerable time, cost, and effort goes into model development and calibration. 4 Prairie Habitat Joint Venture Program John H. Giudice, John Ratti, E. (Oz) Garton Prairie Habitat Joint Venture Program Millions of dollars spent on habitat programs designed to increase annual duck production. need reliable data to evaluate program effectiveness and guide future management and planning. Prairie Habitat Joint Venture Program Brood surveys should furnish simple, direct, and cost-effective data on annual duck production. Most serious problem with brood surveys is visibility bias (i.e., the failure to observe all broods that were present during a survey). Visibility bias is a major cause of inaccuracy in wildlife surveys. Prairie Habitat Joint Venture Program Study Objectives identify and quantify the factors that influence brood visibility; determine the feasibility of using sightability models to estimate brood visibility; and no reliable, timely, and inexpensive methods for estimating annual production of ducks. Duck broods are often secretive and difficult to count. Need for standardized brood-survey procedures that incorporate statistical methods of sampling and sightability correction (sightability models or mark-resight methods) Methods “We used a modified point-count method and 93 radio-marked mallard broods on 7 PHJV Assessment sites (1999-2000) to develop predictive sightability models and assess the relative importance of sightability covariates.” design standardized survey procedures that minimize controllable sources of variation in brood surveys and are compatible with the sightability-model methods. 5 Results Mean visibility of mallard broods was 24%, but detection rates ranged from 13 to 51% depending on year and scale of application Observer experience (years of survey experience) and weather (temperature and wind speed) also influenced detection probability, although effects were less important than percent visual obstruction or brood age Brood visibility was negatively correlated with percent visual obstruction and brood age Results Results Wind speed appeared to be more important than temperature, especially wind speeds >7 km/h. Sightability models developed at the brood and survey-area scale correctly classified 80 and 72% of mallard broods as missed or detected. Sightability Model other examples Anderson, Charles R., Jr. and Lindzey, Frederick G. 1996. Moose sightability model developed from helicopter surveys. Wildl. Soc. Bull. 24:247-259. Ayers, Loren W. and Anderson, Stanley H. 1999. An aerial sightability model for estimating ferruginous hawk population size. J. Wildl. Manage. 63:85-97. Samuel, Michael D.; Garton, Edward O.; Schlegel, Michael W., and Carson, Robert G. 1987. Visibility bias during aerial surveys of elk in northcentral Idaho. J Wildl Manage. 51:622-630. Unsworth, James W.; Kuck, Lonn, and Garton, Edward O. 1990. Elk sightability model validation at the National Bison Range, Montana. Wildl Soc Bull. 18:113-115. Overview of Mark-Recapture Methods Estimation of population size of a geographically and demographically closed but free-ranging population is a common problem encountered by biologists. Overview of Mark-Recapture Methods The earliest approaches to this problem were developed by Petersen in 1896 and later by Lincoln in 1930, where capture-recapture techniques were applied. 6 Simple Mark-Recapture Methods Lincoln-Peterson Method Closed populations, single marking — LincolnPetersen method Lincoln (animal), Petersen (fisheries) Closed populations, multiple markings — Schnabel method Open populations, multiple sampling — JollySeber method. Lincoln-Peterson Method Krebs (1989) provides a succinct method for estimating confidence intervals. If a Petersen population estimate has a very wide confidence interval, you should not place too much faith in it. Jolly-Seber Estimator Jolly-Seber Estimator Jolly-Seber (Jolly 1965) method extends markrecapture to the more biologically realistic situation of open populations. Jolly-Seber Estimator Individuals are marked with tags that are specific to the sampling period. When was this marked individual last captured? The time interval between samples need not be constant, and any number of samples can be accommodated so that series of data extending over may years can be used with this method. 7 Jolly-Seber Estimator Program Mark Program Mark Program Mark Program Mark Program Mark 8 Estimation from mark-resighting surveys Program MARK Program MARK provides parameter estimates from marked animals when they are re-encountered at a later time. Re-encounters can be from dead recoveries (e.g., the animal is harvested), live recaptures (e.g. the animal is re-trapped or resighted), radio tracking, or from some combination of these sources of re-encounters. Check out the description of its capabilities, or information on how to download the program. http://www.cnr.colostate.edu/~gwhite/mark/mark.htm Estimation from mark-resighting surveys Estimation from mark-resighting surveys The mark-resight procedure has been tested with known populations of mule deer (Odocoileus hemionus; Bartmann et al. 1987) and used with white-tailed deer (Rice and Harder 1977), mountain sheep (Furlow et al. 1981, Neal et al. 1993), black grizzly bear (Miller et al. 1987), and coyote (Hein 1992). Alternative Wild Horse and Burro Survey Techniques http://www.fort.usgs.gov/WildHorsePopulations/AlternativeTech.asp USGS Fort Collins Colorado The initial sample of animals is captured and marked with radios, but recaptures of these animals are obtained by observation, not actually recapturing them. The limitation of this procedure is that unmarked animals are not marked on subsequent occasions. The advantage of this procedure is that resightings are generally much cheaper to acquire than physically capturing and handling the animals. Program NOREMARK computes 4 markresight estimators of population abundance, modeling variation of sighting probabilities across time, individual heterogeneity of sighting probabilities, or immigration and emigration from a fixed study area (Eberhardt 1990). Alternative Wild Horse and Burro Survey Techniques population estimates drive nearly all management decisions pertaining to wild horses and burros, accuracy is important. Several widely used techniques exist for conducting aerial population estimates, but they each have limitations. evaluating combinations of these techniques in estimating population numbers of wild horses and burros. 9 Mark-Resight Artificial marking devices, such as radio collars, are considered unacceptable on wild horses. Mark-Resight Simultaneous Double-Count A form of mark-resight Two observers in an aircraft independently observe and record groups of wild horses or wild burros Sighting rates are estimated by comparing sighting records of the two observers Groups seen by both observers must be identified, either by communication with a third observer during the flight, or by matching observations based on time, location, and other characteristics of the sighting Simultaneous Double-Count Simultaneous Double-Count No capture or handling of animals or pre-survey work is required Observer differences estimated and accounted for To ensure that observations are independent (uncorrelated), aerial observers can be isolated through the use of barriers and adherence to strict guidelines and discipline The variable visibility of different groups of animals is a bigger challenge; however, can be addressed through analyses estimate precision can be improved by “marking” or photographing a large portion of the population during the first survey. cost of marking (sighting once) is no higher than for resight: allocating roughly equal effort to the sighting and resighting surveys should produce the most costeffective estimate. Those animals seen by the one observer are the “marked” groups; those that are also seen by the other observer are “resighted.” Sighting probabilities for both observers can be computed from this information Standard mark-resight or Lincoln-Petersen calculations are used to generate a population estimate. Sightability Bias Correction Model The model must hold over space, time, and new observers for estimates to be valid, and populations within each count unit must remain constant during the survey. accounting for multiple sighting factors is also the sightability model’s chief advantage. 10 Distance Sampling Distance Sampling Two observers located on either side of an aircraft record the size of all groups seen and the perpendicular distances from the transect line to each group Transect lines must be placed randomly (or systematically with a random start) but do not necessarily have to be straight; contour lines can be flown instead Location of and distances from the transect line can be measured using GPS and other technologies. Combining Techniques The combination of multiple sources of information overcomes most of the deficiencies of the separate techniques and provides greater power and efficiency For example, the major difficulty in the double-count technique—ensuring similar sighting probabilities for all animals—can be resolved by modeling sightability using covariates in a manner similar to the sightability bias correction model The biggest challenge is likely to be movement of wild horses in response to the approaching aircraft prior to detection by observers. Combining Techniques No pre-calibration of the model is required, and the assumption that the initial calibration applies uniformly over space, time, and observers is eliminated. Similarly, the requirement in distance sampling that all animals near the transect line are spotted can be eliminated by applying data from a double-count to estimate the detection probability on the line Both the mark-resight and distance sampling methods can be made more precise by incorporating sightability covariates. General References General References Jolly, G.M. 1965. Explicit estimates from capture-recapture data with both death and dilution — stochasitc model. Biometrika 52:225247. Seber, G.A.F. 1982. The estimation of animal abundance and related parameters, 2nd ed. Griffin, London. White, Gary C. 1996. NOREMARK: Population estimation from markresighting surveys. Wildlife Society Bulletin. 24:50-52. Krebs, C.J.1989. Ecological Methodology. Harper and Row, New York. Otis, David L.; Burnham, Kenneth P.; White, Gary C., and Anderson, David R. Statistical inference from capture data on closed animal populations. Wildlife Monographs. 1978; 62:1-135. Schnabel, Z.E. 1938. The estimation of the total fish population of a lake. Am. Math. Month. 45:348-352. 11